Anatomy of the endolithic Sonoran Desert lichen Verrucaria rubrocincta Breuss: implications for biodeterioration and biomineralization

Abstract
The anatomy of the endolithic, calcicolous lichen Verrucaria rubrocincta Breuss is described using optical and scanning electron microscopy. This lichen is locally abundant in caliche plates of open desert pavements in the Sonoran Desert of south-western Arizona. The endolithic growth of V. rubrocincta is distinctly layered. The upper layer is a fine-grained calcite (micrite). This layer is sparsely penetrated by hyphae and therefore cannot be interpreted as a lithocortex sensu stricto. Beneath the micrite is the photobiont layer. Below this photobiont layer hyphae form a pseudomedulla penetrating up to 1 cm into the caliche. Calcium oxalates occur in the pseudomedulla but are absent from uninhabited caliche. The analogy of a greenhouse describes the ecophysiological adaptations of this endolithic growth to the environmental extremes of the Sonoran Desert. The micrite acts as a roof and the photobiont layer and pseudomedulla represent the greenhouse interior. Verrucaria rubrocincta has thus evolved a strategy to successfully establish and survive within an extreme environment. Our study illustrates biodeterioration and biomineralization processes acting simultaneously within a single lichen species. Mineralogical evidence suggests that the micrite on the thallus surface is biologically induced. The hyphae of the lichen biodeteriorate the caliche thus forming the pseudomedulla. Simultaneously with this process micrite forms at the surface protecting the thallus from exposure and counter-balancing rock degradation. These combined effects of biodeterioration and biomineralization do not markedly accelerate erosion because inhabited and uninhabited areas of the same plate show similar surface heights.